Coleoptera: Cerambycidae, Buprestidae) Accompanying the Secondary Succession of the Pine Forests of Puszcza Białowieska
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POLSKA AKADEMIA NAUK MUZEUM I INSTYTUT ZOOLOGII FRAGMENTA FAUNISTICA Tom 38 Warszawa, 30 XI 1995 Nr 20 Jerzy M. G u to w s k i Changes in communities of longhorn and buprestid beetles ( Coleoptera: Cerambycidae, Buprestidae) accompanying the secondary succession of the pine forests of Puszcza Białowieska Abstract. Results of a study of Buprestidae and Cerambycidae in the pine forests of Puszcza Białowieska (NE Poland) are presented. 16 and 49 species have been registered respectively. Succession of these taxons in four age classes of forest stand (culture, young stand, pole wood, mature stand) has been analysed, and conspicuous differences have been revealed in species composition, structure of dominance, abundance and diversity. INTRODUCTION Beetles of the families Cerambycidae and Buprestidae are mainly cambio- and xylophagous forms. 189 and 87 species of the respective families have been recorded in Poland to date. The number of species potentially associated with the fresh pine forest habitat and its plants is about 55 and about 25 respectively. Fresh coniferous forests are the dominant habitat type of forest in Poland. The identification of their fauna, including xylo- and cambiophages, is, therefore, important from both the cognitive and practical point of view. Puszcza Białowieska is a unique area of the best preserved forests in the entire European Lowlands. Studies conducted in Puszcza Białowieska make it possible to register facts that are not likely to be detected in other areas, more affected by anthropopression. Owing to this one may investigate relatively untransformed population structures of the species under study, thus serving as a frame of reference for the results of similar studies carried out in other regions of Europe. The aim of this paper was to investigate the secondary succession of the Cerambycidae and Buprestidae fauna associated with forest stands growing in a fresh coniferous forest habitat in Puszcza Białowieska. This is further http://rcin.org.pl 390 J.M. Gutowski associated with the identification of the trophic relations of individual species, their distribution in Puszcza Białowieska, community structure, etc. The groups under study, and particularly Buprestidae, have never before been the subject of extensive succession-centred investigations in fresh pine forests. Papers written to date are concerned predominantly with the entomofauna of pine forests in general, not in terms of forest age classes, or deal with a particular age class, e.g. young stands. Research over the entomofauna of different habitat types of forests, ranking among the first such enterprises in the world, was initiated in Puszcza Białowieska before World War II by J. Karpiński . He explored, among other taxons, Cerambycidae of this region, assigning each species to a forest type he himself had defined (Ka rpiński , 1949). In his study of Pinetum typicum, which may be considered a fresh coniferous forest in the current understanding of this term, he mentioned 21 species. Further analysis of the data provided by Karpiński (op. cit.) has been performed by S z u je c k i (1980). The longhorn beetle fauna of various forest habitat types in Puszcza Niepołomicka has been described by Starzyk (1976), who noted 17 species in a fresh coniferous forest, but did not break the data down into age classes. The figure stands for 21.8% of the total number of species analysed, and, in comparison to 51 species recorded in the fresh mixed forest or 50 registered in the humid forest, is rather low. T omalak (1984), who studied Cerambycidae of the pole wood and mature stands in the environs of Poznań, found 18 species of these beetles in a fresh coniferous forest, which corresponds to 29.5% of the total number of species recorded in the area under investigation. In a similar study of Puszcza Białowieska, G utow ski (1985) found 40 species in fresh coniferous forests, equalling to 36.7% of the total number of species recorded. In comparison to other forest habitat types, the number of species was in the middle range as the greatest number of species (66) has been recorded in a fresh mixed forest and the smallest (11), in a boggy coniferous forest. In his recently published paper on the Roztocze area, G utow ski (1992) has analysed the occurrence of 38 Cerambycidae species of high abundance and frequency in 10 different habitats (including 8 phytosociological groups which are directly referrable to specific forest habitat types). 23 species were found in the subcontinental pine forest (Peucedano-Pinetuni), the dominant forms being: Acanthocimis aedilis, Anastrangalia sanguinolenta, Arhopalus rusticus, Corymbia rubrcc Rhagium inquisitor, Spondylis buprestoides an d Stenurella melanura. Also in Roztocze, this habitat type accommodated a medium number of longhorn beetle species. A description of the entire entomofauna associated with xylem and the subcortical environment in Wielkopolski National Park has been provided by B alazy and M ichalski (1984). The authors recorded 14 Cerambycidae species in the fresh coniferous forest, listing Arhopalus ferus, Pogonocherus decoratus, P. fasciculatus and Prionus coriarius as characteristic of that habitat. Buprestidae found by the authors in fresh coniferous forests included Anthaxia similis, http://rcin.org.pl Cerambycidae, Buprestidae ( Coleoptera) 391 Chalcophora mariana and Phaenops cyanea all the species being characteristic of that habitat. A study of the succession of longhorn beetles depending on the age of the stands has been carried out in Puszcza Niepolomicka by S tarzyk (1977). The study, however, included all the forest habitat types of that region, where fresh coniferous forests constitute only 7.25% of the stand. It cannot be, therefore, determined which Cerambycidae species found, for example, on Scots pine [PiniLS sylvestris L.), were registered in the habitat relevant to us or whether the preferences for trees of particular ages do not depend on the habitat type. It is still interesting to note that there were species which were shown to be affiliated with certain age classes (9 classes: I - 1-20 years, II - 21-40 years, etc). Arhopalus rusticus and Rhagium inquisitor were shown to occur most widely on pine (I-IX age class), while Ergatesfaber occurred only on trees of age class IV and V, and Callidium aeneum - age class III-V. Of the 28 species feeding on pine, 8 were recorded in age class I, 16 in age class II, and then: age class III - 22, IV - 19, V - 15, VI - 10, VII - 10, V III- 5, an d IX - 2. Studies of the succession of selected animal groups in the fresh coniferous forest in Puszcza Białowieska have been conducted by W ołk and G utow ski (1984). Neither Buprestidae n o r Cerambycidae were, however, described separately in the study, which concentrated only on invertebrates as a whole as well as on certain other groups of beetles. There are a number of papers where fragmentary information relevant to the subject of this study may be found. For example, Kinelski and S z u je c k i (1963) have presented the results of studies of secondary pests in young stands of pine (age class I and II) in Puszcza Piska, mentioning 6 species of Cerambycidae (Rhagium inquisitor, Arhopalus rusticus, Callidium sp., M o n o ch a m u s galloprovincialis pistor, Pogonocherus JascicuLatus and Acanthocinus aedilis) and 2 species of Buprestidae (Anthaxia quadripunctata, Chrysobotris igniventris [listed as C. solieri]). W iąckow ski (1957) has examined the entomofauna of pine tree stumps in different stages of wood decomposition (from fresh - right after clear-cutting, to more than 10 years old) also in a fresh coniferous forest habitat. These data may be referred to the first succession stages of the forest stand - clearing and culture, where stumps are the basic source of food for the groups under study. Species associated with stumps at this stage included the longhorn beetle species of Ergates faber and Arhopalus rusticus and the buprestid Chalcophora mariana. B rauns (1975) has listed insects occurring on pine in different age classes. One may suppose (the author did not make it clear) that the study was conducted chiefly in fresh coniferous forests. The following species of longhorn beetles were mentioned as characteristic of tree nurseries, cultures, young stands, pole wood and mature stands: Pogonocherus JascicuLatus (from cultures to mature stands), Tetropium castaneum, T. Juscum, Monochamus sartor (Fabr .), M. sutor and Acanthocinus aedilis (pole wood and mature stands). The only Buprestidae species mentioned w as Phaenops cyanea characteristic of pole wood and mature stands. http://rcin.org.pl 392 J.M. Gutowski Research over the insects of the pine forests in Byelorussia is also presented by Litvinova et al. (1985). There are numerous papers by European authors dealing with longhorn or buprestid beetle communities of various age classes of Scots pine - the main stand-forming species in fresh coniferous forests. Most of them, however, do not contain information on the habitat type where the studies were conducted, which renders them useless from the viewpoint of the present analysis. Scots pine is also abundantly found in the dry, humid and boggy coniferous forest as well as in mixed forests and is also planted in more fertile forest habitats. Comprehensive studies of the entomofauna of fresh coniferous forests, conducted since 1986 in Puszcza Białowieska and other localities by the Institute of Zoology PAS. and the Forest and Wood Protection Institute of Warsaw Agricultural University, have not included the groups discussed in this paper (Bańkow ska 1993). AREAS OF STUDY, MATERIAL, METHODS Areas of study Puszcza Białowieska occupies an area of about 1,500 sq. km. Its western part, belonging to Poland, extends over 624 sq. km. (So kołow ski 1994). Fresh coniferous forests are represented in Puszcza Białowieska by two phytosociological associations (Sokołow ski 1979): Vaccinia vitis-idaeae-Pinetum and Vaccinio myrtiRi-Piceetum. The former, with the stand consisting mainly of Scots pine, is further classified into two subassociations: cladonietosum (a drier one with a high proportion of xerothermal species) and myrtilletosum (slightly more humid with Vaccinium myrtilLus L.